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CONTRIBUTORS Albert H. Banner A. J. Bruce H. Robert Bustard Ann M. Cameron Ailsa M. Clark Robert Endean F. R. Fosberg Barry Goldman Harold Heatwole Jiro Kikkawa Wendell K. Patton Frank Hamilton Talbot BIOLOGY AND GEOLOGY OF CORAL REEFS EDITED BY O. A. JONES Department of Geology University of Queensland St. Lucia, Brisbane Queensland, Australia R. ENDEAN Department of Zoology University of Queensland St. Lucia, Brisbane Queensland, Australia VOLUME III: Biology 2 ACADEMIC PRESS New York San Francisco London A Subsidiary of Harcoutt Brace Jovanooich, Publishers 1976 C O P Y R I G H T © 1976, B Y A C A D E M I C P R E S S , I N C . ALL RIGHTS RESERVED. NO PART O F THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ACADEMIC PRESS, INC. I l l Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 Library of Congress Cataloging in Publication Data Jones, Owen Arthur. Biology and geology of coral reefs. Includes bibliographies. CONTENTS: v. 1. Geology l . - v . 2. Biology 1 . v. 3. Biology 2. 1. Coral reef biology-Collected works. 2. Coral reefs and islands-Collected works. I. Endean, R., joint author. II. Title. QH95.8.J66 574.909'4'2 72-84368 ISBN 0 - 1 2 - 3 8 9 6 0 3 - 7 (v.3) PRINTED IN THE UNITED STATES OF AMERICA To the Great Barrier Reef Committee, its office-bearers and its many members who have worked unremittingly for fifty years to further our knowledge of the Great Barrier Reef; and to the memory of geologist Professor H. C. Richards, one of its founders, and biologist Professor E. J. Goddard, an early enthusiastic supporter LIST OF CONTRIBUTORS Numbers in parentheses indicate the pages on which the authors' contributions begin. H. BANNER, Department of Zoology, University of Hawaii, Honolulu, Hawaii (177) ALBERT A. J. BRUCE, East African Marine Fisheries Research Organization, Mombasa, Kenya, Africa (37) H. Research School of Biological Sciences, Australian National University, Canberra, Australia (343) ROBERT BUSTARD,* ANN M. CAMERON, Department of Zoology, University of Queensland, St. Lucia, Brisbane, Queensland, Australia (155) M. (95) AILSA CLARK, British Museum (Natural History), London, England Department of Zoology, University of Queensland, St. Lucia, Brisbane,. Queensland, Australia (215) ROBERT ENDEAN, F. R. FOSBERG, Botany Department, Natural History Museum, Smithsonian Institution, Washington, D.C. ( 255 ) School of Biological Sciences, Macquarie University, Sydney, Australia (125) BARRY GOLDMAN^ Department of Zoology, University of New England, Armidale, N.S.W., Australia (369) HAROLD HEATWOLE, Department of Zoology, University of Queensland, St Lucia, Brisbane, Queensland, Australia ( 279 ) JIRO KIKKAWA, K. PATTON, Department of Zoology, Ohio Wesleyan University, Delaware, Ohio (1) WENDELL FRANK HAMILTON TALBOT, The Australian Museum, Sydney, Australia (125) * Present address: Airlie Brae, Alyth, Perthshire, Scotland. t Present address: School of Natural Resources, University of the South Pacific, Suva, Fiji. XI GENERAL PREFACE This four-volume work (two volumes covering geological and two biological topics) originated from an article on The Great Barrier Reefs of Australia written by the editors and published in the November 1967 issue of Science Journal. The prime aim of this treatise is to publi